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The calculation of exact reliabilities involving the inversion of mixed model equations poses a heavy computational challenge when the system of equations is large. This has prompted the development of different approximation methods. We give an overview of the various methods and computational approaches in calculating reliability from the era before the animal model to the era of single-step genomic models. The different methods are discussed in terms of modeling, development, and applicability in large dairy cattle populations. The paper also describes the problems faced in reliability computation. Many details dispersed throughout the literature are presented in this paper. It is clear that a universal solution applicable to every model and input data may not be possible, but we point out several efficient and accurate algorithms developed recently for a variety of very large genomic evaluations.

Background The preconditioned conjugate gradient (PCG) method is the current method of choice for iterative solving of genetic evaluations. The relative difference between two successive iterates and the relative residual of the system of equations are usually chosen as a termination criterion for the PCG method in animal breeding. However, our initial analyses showed that these two commonly used termination criteria may report that a PCG method applied to a single-step single nucleotide polymorphism best linear unbiased prediction (ssSNPBLUP) is not converged yet, whereas the solutions are accurate enough for practical use. Therefore, the aim of this study was to propose two termination criteria that have been (partly) developed in other fields, but are new in animal breeding, and to compare their behavior to that of the two termination criteria widely used in animal breeding for the PCG method applied to ssSNPBLUP. The convergence patterns of ssSNPBLUP were also compared to the convergence patterns of single-step genomic BLUP (ssGBLUP). Results Building upon previous work, we propose two termination criteria that take the properties of the system of equations into account. These two termination criteria are directly related to the relative error of the iterates with respect to the true solutions. Based on pig and dairy cattle datasets, we show that the preconditioned coefficient matrices of ssSNPBLUP and ssGBLUP have similar properties when using a second-level preconditioner for ssSNPBLUP. Therefore, the PCG method applied to ssSNPBLUP and ssGBLUP converged similarly based on the relative error of the iterates with respect to the true solutions. This similar convergence behavior between ssSNPBLUP and ssGBLUP was observed for both proposed termination criteria. This was, however, not the case for the termination criterion defined as the relative residual when applied to the dairy cattle evaluations. Conclusion Our results showed that, Numerical Analysis

Background The preconditioned conjugate gradient (PCG) method is the current method of choice for iterative solving of genetic evaluations. The relative difference between two successive iterates and the relative residual of the system of equations are usually chosen as a termination criterion for the PCG method in animal breeding. However, our initial analyses showed that these two commonly used termination criteria may report that a PCG method applied to a single-step single nucleotide polymorphism best linear unbiased prediction (ssSNPBLUP) is not converged yet, whereas the solutions are accurate enough for practical use. Therefore, the aim of this study was to propose two termination criteria that have been (partly) developed in other fields, but are new in animal breeding, and to compare their behavior to that of the two termination criteria widely used in animal breeding for the PCG method applied to ssSNPBLUP. The convergence patterns of ssSNPBLUP were also compared to the convergence patterns of single-step genomic BLUP (ssGBLUP). Results Building upon previous work, we propose two termination criteria that take the properties of the system of equations into account. These two termination criteria are directly related to the relative error of the iterates with respect to the true solutions. Based on pig and dairy cattle datasets, we show that the preconditioned coefficient matrices of ssSNPBLUP and ssGBLUP have similar properties when using a second-level preconditioner for ssSNPBLUP. Therefore, the PCG method applied to ssSNPBLUP and ssGBLUP converged similarly based on the relative error of the iterates with respect to the true solutions. This similar convergence behavior between ssSNPBLUP and ssGBLUP was observed for both proposed termination criteria. This was, however, not the case for the termination criterion defined as the relative residual when applied to the dairy cattle evaluations. Conclusion Our results showed that, Numerical Analysis

Genetic groups, also called unknown or phantom parents groups, are often used in dairy cattle genetic evaluations to account for selection that cannot be accounted for by known genetic relationships. With the advent of genomic evaluations, the theory of genetic groups was extended to the so-called single-step genomic BLUP (ssGBLUP). In short, genetic groups can be fitted in ssGBLUP through regression effects, or by including them in the pedigree and computing the adequate combined pedigree and genomic relationship matrix. In this study, we applied the so-called Quaas and Pollak transformation to a system of equations for single-step SNP BLUP (ssSNPBLUP), such that genetic groups can thereafter be included in the pedigree. The example in this study showed that including the genetic groups in the pedigree for ssSNPBLUP allowed reduced memory burden and computational costs in comparison to genetic groups fitted as covariates.

Background: The preconditioned conjugate gradient (PCG) method is the current method of choice for iterative solving of genetic evaluations. The relative difference between two successive iterates and the relative residual of the system of equations are usually chosen as a termination criterion for the PCG method in animal breeding. However, our initial analyses showed that these two commonly used termination criteria may report that a PCG method applied to a single-step single nucleotide polymorphism best linear unbiased prediction (ssSNPBLUP) is not converged yet, whereas the solutions are accurate enough for practical use. Therefore, the aim of this study was to propose two termination criteria that have been (partly) developed in other fields, but are new in animal breeding, and to compare their behavior to that of the two termination criteria widely used in animal breeding for the PCG method applied to ssSNPBLUP. The convergence patterns of ssSNPBLUP were also compared to the convergence patterns of single-step genomic BLUP (ssGBLUP). Results: Building upon previous work, we propose two termination criteria that take the properties of the system of equations into account. These two termination criteria are directly related to the relative error of the iterates with respect to the true solutions. Based on pig and dairy cattle datasets, we show that the preconditioned coefficient matrices of ssSNPBLUP and ssGBLUP have similar properties when using a second-level preconditioner for ssSNPBLUP. Therefore, the PCG method applied to ssSNPBLUP and ssGBLUP converged similarly based on the relative error of the iterates with respect to the true solutions. This similar convergence behavior between ssSNPBLUP and ssGBLUP was observed for both proposed termination criteria. This was, however, not the case for the termination criterion defined as the relative residual when applied to the dairy cattle evaluations. Conclusion: Our results showed that the PCG method can converge

Background: The single-step single nucleotide polymorphism best linear unbiased prediction (ssSNPBLUP) is one of the single-step evaluations that enable a simultaneous analysis of phenotypic and pedigree information of genotyped and non-genotyped animals with a large number of genotypes. The aim of this study was to develop and illustrate several computational strategies to efficiently solve different ssSNPBLUP systems with a large number of genotypes on current computers. Results: The different developed strategies were based on simplified computations of some terms of the preconditioner, and on splitting the coefficient matrix of the different ssSNPBLUP systems into multiple parts to perform its multiplication by a vector more efficiently. Some matrices were computed explicitly and stored in memory (e.g. the inverse of the pedigree relationship matrix), or were stored using a compressed form (e.g. the Plink 1 binary form for the genotype matrix), to permit the use of efficient parallel procedures while limiting the required amount of memory. The developed strategies were tested on a bivariate genetic evaluation for livability of calves for the Netherlands and the Flemish region in Belgium. There were 29,885,286 animals in the pedigree, 25,184,654 calf records, and 131,189 genotyped animals. The ssSNPBLUP system required around 18 GB Random Access Memory and 12 h to be solved with the most performing implementation. Conclusions: Based on our proposed approaches and results, we showed that ssSNPBLUP provides a feasible approach in terms of memory and time requirements to estimate genomic breeding values using current computers.

Background: The preconditioned conjugate gradient (PCG) method is an iterative solver of linear equations systems commonly used in animal breeding. However, the PCG method has been shown to encounter convergence issues when applied to single-step single nucleotide polymorphism BLUP (ssSNPBLUP) models. Recently, we proposed a deflated PCG (DPCG) method for solving ssSNPBLUP efficiently. The DPCG method introduces a second-level preconditioner that annihilates the effect of the largest unfavourable eigenvalues of the ssSNPBLUP preconditioned coefficient matrix on the convergence of the iterative solver. While it solves the convergence issues of ssSNPBLUP, the DPCG method requires substantial additional computations, in comparison to the PCG method. Accordingly, the aim of this study was to develop a second-level preconditioner that decreases the largest eigenvalues of the ssSNPBLUP preconditioned coefficient matrix at a lower cost than the DPCG method, in addition to comparing its performance to the (D)PCG methods applied to two different ssSNPBLUP models. Results: Based on the properties of the ssSNPBLUP preconditioned coefficient matrix, we proposed a second-level diagonal preconditioner that decreases the largest eigenvalues of the ssSNPBLUP preconditioned coefficient matrix under some conditions. This proposed second-level preconditioner is easy to implement in current software and does not result in additional computing costs as it can be combined with the commonly used (block-)diagonal preconditioner. Tested on two different datasets and with two different ssSNPBLUP models, the second-level diagonal preconditioner led to a decrease of the largest eigenvalues and the condition number of the preconditioned coefficient matrices. It resulted in an improvement of the convergence pattern of the iterative solver. For the largest dataset, the convergence of the PCG method with the proposed second-level diagonal preconditioner was slower than the DPCG method, but it perf

Background The preconditioned conjugate gradient (PCG) method is an iterative solver of linear equations systems commonly used in animal breeding. However, the PCG method has been shown to encounter convergence issues when applied to single-step single nucleotide polymorphism BLUP (ssSNPBLUP) models. Recently, we proposed a deflated PCG (DPCG) method for solving ssSNPBLUP efficiently. The DPCG method introduces a second-level preconditioner that annihilates the effect of the largest unfavourable eigenvalues of the ssSNPBLUP preconditioned coefficient matrix on the convergence of the iterative solver. While it solves the convergence issues of ssSNPBLUP, the DPCG method requires substantial additional computations, in comparison to the PCG method. Accordingly, the aim of this study was to develop a second-level preconditioner that decreases the largest eigenvalues of the ssSNPBLUP preconditioned coefficient matrix at a lower cost than the DPCG method, in addition to comparing its performance to the (D)PCG methods applied to two different ssSNPBLUP models. Results Based on the properties of the ssSNPBLUP preconditioned coefficient matrix, we proposed a second-level diagonal preconditioner that decreases the largest eigenvalues of the ssSNPBLUP preconditioned coefficient matrix under some conditions. This proposed second-level preconditioner is easy to implement in current software and does not result in additional computing costs as it can be combined with the commonly used (block-)diagonal preconditioner. Tested on two different datasets and with two different ssSNPBLUP models, the second-level diagonal preconditioner led to a decrease of the largest eigenvalues and the condition number of the preconditioned coefficient matrices. It resulted in an improvement of the convergence pattern of the iterative solver. For the largest dataset, the convergence of the PCG method with the proposed second-level diagonal preconditioner was slower than the DPCG meth, green, Numerical Analysis

Background: The single-step single nucleotide polymorphism best linear unbiased prediction (ssSNPBLUP) method, such as single-step genomic BLUP (ssGBLUP), simultaneously analyses phenotypic, pedigree, and genomic informa- tion of genotyped and non-genotyped animals. In contrast to ssGBLUP, SNP effects are fitted explicitly as random effects in the ssSNPBLUP model. Similarly, principal components associated with the genomic information can be fitted explicitly as random effects in a single-step principal component BLUP (ssPCBLUP) model to remove noise in genomic information. Single-step genomic BLUP is solved efficiently by using the preconditioned conjugate gradi- ent (PCG) method. Unfortunately, convergence issues have been reported when solving ssSNPBLUP by using PCG. Poor convergence may be linked with poor spectral condition numbers of the preconditioned coefficient matrices of ssSNPBLUP. These condition numbers, and thus convergence, could be improved through the deflated PCG (DPCG) method, which is a two-level PCG method for ill-conditioned linear systems. Therefore, the first aim of this study was to compare the properties of the preconditioned coefficient matrices of ssGBLUP and ssSNPBLUP, and to document convergence patterns that are obtained with the PCG method. The second aim was to implement and test the effi- ciency of a DPCG method for solving ssSNPBLUP and ssPCBLUP. Results:For two dairy cattle datasets, the smallest eigenvalues obtained for ssSNPBLUP (ssPCBLUP) and ssGBLUP, both solved with the PCG method, were similar. However, the largest eigenvalues obtained for ssSNPBLUP and ssPCB- LUP were larger than those for ssGBLUP, which resulted in larger condition numbers and in slow convergence for both systems solved by the PCG method. Different implementations of the DPCG method led to smaller condition num- bers, and faster convergence for ssSNPBLUP and for ssPCBLUP, by deflating the largest unfavourable, green, Numerical Analysis

Background: The single-step single nucleotide polymorphism best linear unbiased prediction (ssSNPBLUP) method, such as single-step genomic BLUP (ssGBLUP), simultaneously analyses phenotypic, pedigree, and genomic informa- tion of genotyped and non-genotyped animals. In contrast to ssGBLUP, SNP effects are fitted explicitly as random effects in the ssSNPBLUP model. Similarly, principal components associated with the genomic information can be fitted explicitly as random effects in a single-step principal component BLUP (ssPCBLUP) model to remove noise in genomic information. Single-step genomic BLUP is solved efficiently by using the preconditioned conjugate gradi- ent (PCG) method. Unfortunately, convergence issues have been reported when solving ssSNPBLUP by using PCG. Poor convergence may be linked with poor spectral condition numbers of the preconditioned coefficient matrices of ssSNPBLUP. These condition numbers, and thus convergence, could be improved through the deflated PCG (DPCG) method, which is a two-level PCG method for ill-conditioned linear systems. Therefore, the first aim of this study was to compare the properties of the preconditioned coefficient matrices of ssGBLUP and ssSNPBLUP, and to document convergence patterns that are obtained with the PCG method. The second aim was to implement and test the effi- ciency of a DPCG method for solving ssSNPBLUP and ssPCBLUP. Results:For two dairy cattle datasets, the smallest eigenvalues obtained for ssSNPBLUP (ssPCBLUP) and ssGBLUP, both solved with the PCG method, were similar. However, the largest eigenvalues obtained for ssSNPBLUP and ssPCB- LUP were larger than those for ssGBLUP, which resulted in larger condition numbers and in slow convergence for both systems solved by the PCG method. Different implementations of the DPCG method led to smaller condition num- bers, and faster convergence for ssSNPBLUP and for ssPCBLUP, by deflating the largest unfavourable, green, Numerical Analysis

Background: The single-step single nucleotide polymorphism best linear unbiased prediction (ssSNPBLUP) method, such as single-step genomic BLUP (ssGBLUP), simultaneously analyses phenotypic, pedigree, and genomic information of genotyped and non-genotyped animals. In contrast to ssGBLUP, SNP effects are fitted explicitly as random effects in the ssSNPBLUP model. Similarly, principal components associated with the genomic information can be fitted explicitly as random effects in a single-step principal component BLUP (ssPCBLUP) model to remove noise in genomic information. Single-step genomic BLUP is solved efficiently by using the preconditioned conjugate gradient (PCG) method. Unfortunately, convergence issues have been reported when solving ssSNPBLUP by using PCG. Poor convergence may be linked with poor spectral condition numbers of the preconditioned coefficient matrices of ssSNPBLUP. These condition numbers, and thus convergence, could be improved through the deflated PCG (DPCG) method, which is a two-level PCG method for ill-conditioned linear systems. Therefore, the first aim of this study was to compare the properties of the preconditioned coefficient matrices of ssGBLUP and ssSNPBLUP, and to document convergence patterns that are obtained with the PCG method. The second aim was to implement and test the efficiency of a DPCG method for solving ssSNPBLUP and ssPCBLUP. Results: For two dairy cattle datasets, the smallest eigenvalues obtained for ssSNPBLUP (ssPCBLUP) and ssGBLUP, both solved with the PCG method, were similar. However, the largest eigenvalues obtained for ssSNPBLUP and ssPCBLUP were larger than those for ssGBLUP, which resulted in larger condition numbers and in slow convergence for both systems solved by the PCG method. Different implementations of the DPCG method led to smaller condition numbers, and faster convergence for ssSNPBLUP and for ssPCBLUP, by deflating the largest unfavourable eigenvalues. Conclusions: Poor convergence of ssSNPBLUP